CAUSE SPECIFIC MORTALITY AND ANTI-PREDATOR BEHAVIOR IN MIDWESTERN SONGBIRDS A Dissertation Presented to The Faculty of the Graduate School At the University of Missouri In Partial Fulfillment Of the Requirements for the Degree Doctor of Philosophy By WILLIAM ANDREW COX John Faaborg, Dissertation Supervisor JULY 2011 © Copyright by William Andrew Cox 2011 All Rights Reserved The undersigned, appointed by the dean of the Graduate School, have examined the Dissertation entitled CAUSE SPECIFIC MORTALITY AND ANTI-PREDATOR BEHAVIOR IN MIDWESTERN SONGBIRDS Presented by William Andrew Cox A candidate for the degree of Doctor of Philosophy And hereby certify that, in their opinion, it is worthy of acceptance. Professor John Faaborg Professor Frank R. Thompson III Professor Ray Semlitsch Professor Lori Eggert ACKNOWLEDGEMENTS The number of people I need thank for love, patience, and guidance throughout the design, execution, and writing of this dissertation is constrained only by the time I spend contemplating this page. I relied upon the unerring support of those closest to me. I depended on the kindness of strangers. I was given relentless effort in the field from a number of extraordinary people. There are simply too many people to thank and too few words to do them justice. I mean it. Nevertheless, I’ll do my best. First and foremost, family. My father and sisters have been nothing but supportive despite my rather poor prioritization of what matters most these past six years. I thank them for their patience. I also thank my sisters for Rachel, Maddie, and Bea. Such beautiful girls! Bob and Sandy Simpler have been unbelievably generous since the day I met them 15 years ago. Their patience and help has been invaluable; a prototype battery backpack was Bob’s construct and saved me from certain death during my pilot field season. And thanks to Michael, Gemma, Noah and Tia for housing and feeding us on our trips out east. Never was there an easier place to visit and briefly forget about work. When returned to the University of Florida in 2003 for a year of post- baccalaureate study, I didn’t know anything about anything. I am grateful to Doug Levey for being such an inspirational teacher and Jeremy Kirchman for his encouragement and friendship. I especially thank Rebecca Kimball and Ed Braun for inviting me into their lab, introducing me to the nuts and bolts of the ii scientific process, and putting me on two publications that paved the way for much that followed. I thank them for making all the hard work so much fun and for their continued friendship. Tom Martin and his crew (especially T.J. Fontaine) deserve special thanks for taking me to some of the world’s most beautiful places and for inspiring me time and again to work and think hard. Frank Thompson, Ray Semlitsch, and Lori Eggert have all helped me immeasurably over the past six years. I am fortunate beyond words to have a committee stacked with such caring, bright people. They have been excellent mentors in work and in life; I love them all. John welcomed me into his lab and house as if I were family. I appreciate all that he has done for me, and thank him and Janice for taking care of me throughout my time in Columbia. The Faaborg- led trips to Mexico and Puerto Rico will forever be etched in my heart and inspired me to work to share similar experiences with students in the future. Other MU staff and faculty who I owe a debt of gratitude to (in no particular order): Josh Millspaugh, Jim Carrel, Nila Emerich, Alan Marshall, Josh Hartley, Tyeece Little. My fellow graduate students have been an invaluable source of academic and psychological support. There are many current and former avian ecology lab members who deserve my thanks. The big three are Robin Hirsch-Jacobson (and family!), Judith Toms, and Cara Joos, all of whom have been invaluable colleagues and friends. As have Rafael Brito-Aguilar, Alicia Burke, Andy George, Chris Merkord, Marissa Ahlering, Ernesto Ruelas Inzunzas, Jen White, Chad Rittenhouse, Shane Pruett, Conor McGowan, Sarah Wolken, and Kaylan iii Kemink. Bird people rock. Non-bird people such as Bill Peterman, Jen Hamel, Stephanie Schuttler and many others have also been great. A project such as this cannot succeed without dedicated, fanatically hard- working field assistants. I thank everyone who helped, but especially the following: Riccardo Ton, master nest searcher, cook, humorist. Kelly Schaeffer, Acadian queen in 2009 and all-around machine in 2010. And Curtis Kukal, world’s nicest guy and diligent field tech. I know I learned more from him than he from me. Finally, I want to thank my immediate family. Rabbit, Ike, and Theta have all been instant stress relievers. I saved the best for last; my wife Allison has done it all over the past six years. She is kind, bright, patient, diligent, funny, and sympathetic, all of which she used to improve the quality of my work and life. Lou Gehrig was wrong – I am the luckiest guy on the planet. iv TABLE OF CONTENTS ACKNOWLEDGEMENTS .................................................................................... ii LIST OF TABLES .............................................................................................. viii LIST OF FIGURES ............................................................................................... x ABSTRACT ........................................................................................................ xiii CHAPTER 1. PREDATOR-SPECIFIC RATES OF PREDATION EXPLAIN VARIATION IN NEST SURVIVAL ........................................................................................ 1 ABSTRACT ...................................................................................................... 1 INTRODUCTION .............................................................................................. 2 METHODS ....................................................................................................... 5 Study sites and focal species ....................................................................... 5 Nest monitoring and camera placement ....................................................... 6 Covariates .................................................................................................... 8 Analysis ........................................................................................................ 9 Models ........................................................................................................ 10 RESULTS ....................................................................................................... 12 DISCUSSION ................................................................................................. 14 ACKNOWLEDGEMENTS ............................................................................... 20 LITERATURE CITED ..................................................................................... 22 2. LANDSCAPE AND NEST SITE FACTORS INFLUENCE PREDATOR- SPECIFIC RATES OF NEST PREDATION ................................................... 39 ABSTRACT .................................................................................................... 39 INTRODUCTION ............................................................................................ 40 v METHODS ..................................................................................................... 43 Data collection ............................................................................................ 43 Analysis ...................................................................................................... 45 RESULTS ....................................................................................................... 48 DISCUSSION ................................................................................................. 49 ACKNOWLEDGEMENTS ............................................................................... 55 LITERATURE CITED ..................................................................................... 57 3. NEST VISITATION RATES VARY IN RESPONSE TO PREDATION RISK FROM A DIVERSE SUITE OF PREDATORS ................................................ 69 ABSTRACT .................................................................................................... 69 INTRODUCTION ............................................................................................ 70 METHODS ..................................................................................................... 73 Data collection ............................................................................................. 73 Analysis ....................................................................................................... 75 RESULTS ....................................................................................................... 77 DISCUSSION ................................................................................................. 79 ACKNOWLEDGMENTS ................................................................................. 83 LITERATURE CITED ..................................................................................... 84 4. DEVELOPMENT OF CAMERA TECHNOLOGY FOR MONITORING NESTS ........................................................................................................... 95 ABSTRACT .................................................................................................... 95 INTRODUCTION ...........................................................................................